BRIGHT RED & YELLOW BROMELIAD FLOWERS

Aechmea tillandsioides

Biotopo Chocón Machacas Livingston, Izabal

NICHOLAS HELLMUTH BRIGHT RED & YELLOW BROMELIAD FLOWERS

Aechmea tillandsioides

Biotopo Chocón Machacas Livingston, Izabal

NOVEMBER 2020 CREDITS APPRECIATION

The helpful individuals listed below are all part of Assistance for local Access, the FLAAR Mesoamerica research and field work Municipio de Livingston Daniel Esaú Pinto Peña, Alcalde of Livingston team. The office research team, webmaster, and (Izabal, ). web designers are additional individuals in the main office in Guatemala City. Since each report Initiation of the Project of Cooperation, is a different or animal, the individuals who February and March 2020 assist in preparing the bibliography, Edwin Mármol Quiñonez, Coordinación de identification and botanical information category Cooperación de Livingston (Izabal, Guatemala) are, not the same for each report. Assistance for Knowledge of Author of Alta Verapaz “Don Chuz”, Fabio Jesus Ical Xo Nicholas Hellmuth

Assistance for Knowledge of plants of Péten Botanical Information Compiler “Teco”, Moises Daniel Pérez Diaz, Nicholas Hellmuth park ranger, PNYNN Chloe Alejandra Valenzuela Assistance for Knowledge of plants at Tapón Creek and Taponcito Creek FUNDAECO Species Identification Team Lucas Cuz Victor Mendoza Lancheros from Muelle Municipal to Field Trip Base Senaida Ba Camp & Back Omar Suchite Keneth William De La Cruz Bibliography Team Nicholas Hellmuth Lancheros and Guide Buena Vista Vivian Hurtado Tapon Creek Edgar Alexander Cuz Choc

Editor Vivian Díaz MUNICIPALIDAD DE Livingston Photographers Nicholas Hellmuth ______María Alejandra Gutiérrez PHOTO FROM FRONT COVER David Arrivillaga Aechmea tillandsioides

Photo by: Vivian Díaz, FLAAR Mesoamerica, Photography Aassistants Feb. 14, 2020. Livingston, Izabal, Guatemala. Senaida Ba Mucu Camera: Google Pixel 3 XL.

Manager of Design and Layout PHOTO FROM TITLE PAGE Andrea Sánchez Díaz Aechmea tillandsioides

Photo by: Vivian Díaz, FLAAR Mesoamerica, Layout of this English Edition Feb. 14, 2020. Livingston, Izabal, Guatemala. Cristina Ríos Camera: Google Pixel 3 XL. CONTENTS In what Ecosystems can you find native Aechmea tillandsioides? 1 FLAAR Mesoamérica INTRODUCTION TO BROMELIADS IN GUATEMALA

Bromeliad flowers are popular around the world. Every major botanical garden is filled with bromeliads and orchids. There are bromeliad societies in many major cities.

In Guatemala your eyes will be overwhelmed by the flowering bromeliads, orchids, and vines in the humid mountain areas of Alta Verapaz. We at FLAAR Mesoamérica have driven the back roads of Alta Verapaz with 4-wheel drive, and hiked to areas not even reachable by 4WD, so have experienced lots of bromeliads in Alta Verapaz.

Aechmea tillandsioides Baker.

Photo by: Victor Castillo, FLAAR Mesoamerica, Feb. 14, 2020. Río Chocón Machacas, El Golfete, Livingston, Izabal, Guatemala.

Camera: Sony Alpha A9 II. Lens: Sony FE 200-600mm G OSS. Settings: 1/640 sec; f/6.3; ISO 2,000. 2 FLAAR Mesoamérica

At Parque Nacional Yaxha, Nakum and Naranjo, coordinated by the park administrators of IDAEH and CONAP, and with the assistance of the park rangers, we also learned a lot about bromeliads there in Petén. Although of course I like epiphytic bromeliads, I experienced a lot of terrestrial bromeliads as well: park ranger “Teco” (Moises Daniel Pérez Díaz) took us to special ecosystems where we photographed Aechmea magdalenae often in masses of several hundred surrounding a seasonal “aguada” (aguada means an area which often inundates on the rainy season).

The biodiversity of Municipio de Livingston is another paradise of bromeliads. Since you can take boat trips along the shores of Rio Dulce, El Golfete, and Canyon Rio Dulce, you can see the bromeliads flowering up in the trees. And when you hike the trails of the several nature reserves in this Caribbean area, you see even more bromeliads. We recently went up every single river on the north side of El Golfete: several had awesome paradise of bromeliads, many in flower (November 2020).

With the present FLAAR report, I would like to introduce one of the bromeliads we saw most often during mid-March Exploring Livingston. 2020: Aechmea tillandsioides. Photo by: Juan Pablo Fumagalli, FLAAR Mesoamerica, Mar. 9, 2020. Livingston, Izabal.

Camera: Google Pixel 3 XL. 3 FLAAR Mesoamérica

Aechmea tillandsioides. Family . “Piñuela” is one of the common names given in some areas of Guatemala. Photo by: Nicholas Hellmuth, FLAAR Mesoamerica, Mar. 11, 2020. Chocón Machacas, Livingston, Izabal. Camera: Google Pixel 3 XL.

If you want to discover more from Parque Nacional Yaxha, Nakum and Naranjo visit our site: https://flaar-mesoamerica.org/projects- national-park-yaxha-nakum-naranjo/ 4 FLAAR Mesoamérica

Aechmea tillandsioides Baker.

Photo by: Nicholas Hellmuth, FLAAR Mesoamerica, Nov. 8, 2020. Río Lámpara, Livingston. Camera: iPhone 11 Pro Max. 5 FLAAR Mesoamérica FULL BOTANICAL NAME AND HABIT

Aechmea tillandsioides (Mart. ex Schult. & Schult.f.) Baker

Habit: herb, epiphytic. This bromeliad species survives happily “on the ground” if the branch holding it also falls down. However, it occasionally survives by itself on the ground even without the branch. Nonetheless, it is definitely an arboreal plant.

Aechmea tillandsioides. Family Bromeliaceae. “Piñuela” is one of the common names given in some areas of Guatemala.

Photo by: David Arrivillaga, FLAAR Mesoamerica, Mar. 11, 2020. Municipio de Livingston, Izabal. Camera: Nikon D810. Lens: Nikon 200mm AF-D Tele-Macro. Settings: 1/250 sec; f/14; ISO 3,200. 6 FLAAR Mesoamérica

We found it in full bloom in February and still in March in Biotope Chocón Machacas, El Golfete, Municipio de Livingston.

Experienced botanist of MOBOT, Dr Thomas Croat, found it at Yaxha (ironically near the FLAAR camp in 1973; FLAAR was founded in 1969 to map Yaxha and we worked there from 1970 through 1974).

Only two specimens are in herbaria on-line for all of Izabal: so in two short field trips the team of FLAAR Mesoamerica found as many individual plants as botanists from USA have collected in the past half century.

Aechmea tillandsioides Baker. Family Bromeliaceae.

Photo by: Nicholas Hellmuth, FLAAR Mesoamerica, Jul. 9, 2019. Naranjo, Yaxha, Petén, Guatemala. Camera: iPhone XS. Aechmea tillandsioides. Family Bromeliaceae. “Piñuela” is one of the common names given in some areas of Guatemala.

Photo by: Vivian Díaz, FLAAR Mesoamerica, Feb. 14, 2020. Livingston, Izabal. Camera: Google Pixel 3 XL. 8 FLAAR Mesoamérica

LOCAL NAMES FOR AECHMEA TILLANDSIOIDES

Surely this plant has local names but most publications do not provide any list. We will ask when we return to Livingston Municipality in a future field trip. In the meantime, two local names are provided by Standley and Steyermark: Pie de gallo (Petén); eck (Alta Verapaz) (1958: 386).

Aechmea tillandsioides, a bromeliad flower coming out of the treetop.

Photo by: Victor Mendoza, FLAAR Mesoamerica, Nov. 8, 2020. Río Tatín, Río Dulce, Municipio de Livingston. Camera: Sony RX10 IV. Lens: Sony FE 200-600mm G OSS. Settings: 1/250 sec; f/4; ISO 200. 9 FLAAR Mesoamérica

SYNONYMS FOR AECHMEA TILLANDSIOIDES

• Aechmea chiriquensis Baker • Aechmea kienastii E.Morren ex Mez • Aechmea tillandsioides var. kienastii (E. Morren ex Mez) L.B.Sm. • Aechmea tillandsioides var. tillandsioides • Aechmea vrieseoides Baker • Aechmea xiphophylla Baker • Billbergia gracilis Poepp. ex Beer • Billbergia tillandsioides Mart. ex Schult. & Schult.f. • Ortgiesia tillandsioides (Mart. ex Schult. & Schult.f.) Regel • Ortgiesia tillandsioides var. nidulans Regel • Platyaechmea tillandsioides (Mart. ex Schult. & Schult.f.) L.B.Sm. & W.J.Kress • Platyaechmea tillandsioides var. kienastii (E. Morren ex Mez) L.B.Sm. & W.J.Kress • Platystachys gracilis Beer [Invalid] Synonym • Portea tillandsioides (Mart. ex Schult. & Schult.f.) G.Nicholson

Aechmea tillandsioides Baker.

Photo by: Senaida Ba, FLAAR Mesoamerica, Nov. 8, 2020. Río Lámpara, Río Dulce, Livingston.

Camera: Nikon D5. Lens: Nikon AF-S NIKKOR 400mm FL ED VR. Settings: 1/160 sec; f/8; ISO 800. Aechmea tillandsioides. Family Bromeliaceae. “Piñuela” is one of the common names given in some areas of Guatemala. Photo by: Victor Castillo, FLAAR Mesoamerica, Nov. 7, 2020. Chocón Machacas, Livingston, Izabal. Camera: Sony Alpha A9 II. Lens: Sony FE 200-600mm G OSS. Settings: 1/640 sec; f/6.3; ISO 2,ooo.

Aechmea tillandsioides Baker. Family Bromeliaceae. Photo by: María Alejandra Gutierrez, FLAAR Mesoamerica, Nov. 7, 2020. Chocón Machacas, Livingston, Izabal. Camera: Sony Alpha A9 II. Lens: Sony FE 200-600mm G OSS. Settings: 1/1,000 sec; f/10; ISO 6,400. 11 FLAAR Mesoamérica CLOSE RELATIVES OF AECHMEA TILLANDSIOIDES PRESENT IN GUATEMALA

Aechmea bracteata But Aechmea tessmanii is from South Aechmea bromeliifolia America, not , and is a Aechmea lueddemanniana different species. Aechmea tessmanii is in Aechmea magdalenae fact an accepted name for this different Aechmea mexicana species; so, it may not be considered a Aechmea tessmannii (if you are not a synonym of Aechmea tillandsioides. botanist with a magnifying glass), looks a bit similar to Aechmea tillandsioides.

Aechmea bracteata Griseb.

Photo by: Nicholas Hellmuth, FLAAR Mesoamerica, Jun. 3, 2019. Yaxha, Petén, Guatemala. Camera: iPhone XS. 12 FLAAR Mesoamérica

Aechmea bracteata. This bromeliad is known as “gallito” in some Spanish speaking countries.

Photo by: Nicholas Hellmuth, FLAAR Mesoamerica, Nov. 8, 2020. Yaxha, Petén. Camera: Nikon D810. Lens: Nikon 200mm AF-D Tele-Macro. Settings: 1/200 sec; f/1o; ISO 3,200. 13 FLAAR Mesoamérica AECHMEA TILLANDSIOIDES HAS BEEN COLLECTED AT PARQUE NACIONAL TIKAL

• Missouri Botanical Garden • Catalog #: 1793184 • Taxon: Aechmea tillandsioides var. kienastii (E. Morren ex Mez) L.B. Sm. • Family: Bromeliaceae • Determiner: L.B. Smith, • Collector: C.L. Lundell 15498 • Date: 1959-01-29 • Locality: Guatemala, Petén, Tikal National Park, Tikal. In ramonal covering the ruins 17.23 -89.61

Aechmea tillandsioides. Family Bromeliaceae. “Piñuela” is one of the common names given in some areas of Guatemala.

Photo by: Nicholas Hellmuth, FLAAR Mesoamerica, Nov. 8, 2020. Livingston, Izabal. Camera: Nikon D810. Lens: Nikon 200mm AF-D Tele-Macro. Settings: 1/320 sec; f/10; ISO 800. 14 FLAAR Mesoamérica

SAME FOR YAXHA, NAKUM AND NARANJO, AECHMEA TILLANDSIOIDES HAS BEEN COLLECTED AT YAXHA?

• Missouri Botanical Garden • Catalog #: 1976048 • Taxon: Aechmea tillandsioides (Mart. ex Schult. & Schult. f.) Baker • Family: Bromeliaceae • Determiner: J. Utley • Collector: Thomas B. Croat 24678 • Date: 1973-06-18 • Locality: Guatemala, Petén, Vicinity of Archeological camp on north shore of Lake Yaxha. Disturbed forest 17.06 -89.39 • Elevation: 152 meters (499ft)

• Missouri Botanical Garden • Catalog #: 100276492 • Taxon: Aechmea tillandsioides (Mart. ex Schult. & Schult. f.) Baker • Family: Bromeliaceae • Determiner: M. Véliz • Collector: 1265 • Date: 1989-03-10 • Locality: Guatemala, Petén, Yaxha, El Petén.

Guzmania scherzeriana, “Gallinzaco” is the common name given on Livingston for this Bromeliad.

Photo by: Nicholas Hellmuth, FLAAR Mesoamerica, Nov. 10, 2020. Mirador El Cañón, El Golfete, Río Negro, Livingston.

Camera: Nikon D810. Lens: Nikon 200mm AF-D Tele-Macro. Settings: 1/250 sec; f/11; ISO 2,560. Aechmea tillandsioides Baker. Photo by: Victor Mendoza, FLAAR Mesoamerica, Nov. 8, 2020. Río Tatin, Río Dulce, Izabal. Camera: Sony RX10 IV. Lens: Sony FE 90mm Macro G OSS. Settings: 1/80 sec; f/4; ISO 8oo.

Aechmea tillandsioides. Family Bromeliaceae. “Piñuela” is one of the common names given in some areas of Guatemala. Photo by: Nicholas Hellmuth, FLAAR Mesoamerica, Oct. 28, 2018. Yaxha, Petén. Camera: Nikon D810. Lens: Nikon 200mm AF-D Tele-Macro. Settings: 1/250 sec; f/13; ISO 640. 16 FLAAR Mesoamérica

Vilma Fialko has found Aechmea tillandsioides at Naranjo area of the park. The team of FLAAR Mesoamérica has noticed Aechmea tillandsioides at the Yaxha and Nakum areas of this extensive park.

Aechmea tillandsioides Baker.

Photo by: Nicholas Hellmuth, FLAAR Mesoamerica, Nov. 20, 2020. Mirador El Cañón, El Golfete, Río Negro, Livingston. Camera: Nikon D810. Lens: Nikon 200mm AF-D Tele-Macro. Settings: 1/250 sec; f/9; ISO 4,000. 17 FLAAR Mesoamérica

BRIEF MENTION OF AECHMEA TILLANDSIOIDES BY STANDLEY AND RECORD (1936)

Only one single solitary line on this gorgeous plant. No botanical description; only a mention in passing.

Aechmea tillandsioides Baker. Big Creek, Schipp 98.

(Standley and Record 1936: 90).

Biotopo Chocón Machacas

Photo by: Victor Mendoza, FLAAR Mesoamerica, Feb. 14, 2020. Biotopo Chocón Machacas, Livingston, Izabal. Camera: Google Pixel 3 XL. 18 FLAAR Mesoamérica

BOTANICAL DESCRIPTION OF AECHMEA TILLANDSIOIDES BY STANDLEY AND STEYERMARK (1958)

Aechmea tillandsioides (Mart.) Baker var. Kienastii (E. Morr. ex Mez) L. B. Smith, Caldasia 5: 5. 1942. Ae. Kienastii E. Morr. ex Mez in DC. Monog. Phan. 9: 243. 1896. Ae. squarrosa Baker, Jour. Bot. 28: 305. 1890 (type from Boca de Polochic, Dept. Izabal, J. D. Smith 1823), not Baker, 1889. Pie de gallo (Petén); eck (Alta Verapaz). Figure 66. Epiphytic in forests, up to 300 meters; Petén; Alta Verapaz; Izabal. Southern México; Central America; ; Amazon Basin. The typical variety is in Colombia, the Amazon Basin and Guiana. Leaves 5-9 dm. long with serrate blades 10-65 mm. wide; scape short, its bracts remote, lanceolate, red, serrate; inflorescence simple or digitate (pinnate in the typical variety), white-floccose at first; spikes oblong, 4-11 cm. long; rhachis excavated; floral bracts distichous, imbricate, 10-17 mm. long, convex, nerved; sepals asymmetric, 7-10 mm. long; petals dark purple.

(Standley and Steyermark 1958: 386-387)

Aechmea tillandsioides. Family Bromeliaceae. “Piñuela” is one of the common names given in some areas of Guatemala.

Photo by: David Arrivillaga, FLAAR Mesoamerica, Feb. 14, 2020. Biotopo Chocón Machacas, Municipio de Livingston. Camera: Sony Alpha A9 II. Lens: Sony FE 90mm Macro G OSS. Settings: 1/250 sec; f/9; ISO 640. Aechmea tillandsioides. Family Bromeliaceae.

Photo by: David Arrivillaga, FLAAR Mesoamerica, Feb. 14, 2020. Biotopo Chocón Machacas, Municipio de Livingston. Camera: Sony Alpha A9 II. Lens: Sony FE 90mm Macro G OSS. Settings: 1/250 sec; f/9; ISO 640. 20 FLAAR Mesoamérica COVERAGE OF AECHMEA TILLANDSIOIDES IN OTHER PERTINENT BOOKS

Harvard University Herbaria Aechmea tillandsioides (Mart. & Schult, f.) Baker var. tillandsioides , J. Bot. 17: 134. 1879. Basionym: Billbergia tillandsioides Mart. & Schult, f., Syst. Veg. 7, 2: 1269. 1830. Synonym: Platyaechmea tillandsioides (Mart. & Schult, f.) L. B. Sm. & Kress var tillandsioides, Phytologia 69(4): 274. 1990. . Campeche: Calakmul, E. Martinez et al. 27 828- A (MEXU); Quintana Roo: Selva Mediana, 1 km E of Chanká Veracruz, selva mediana con Brosimum , Pithecellobium y Vitex, epifita acaulescente con inflorescencias rojas, 7 Mayo 1983, E . Cabrera and R. Durán 4668 (CIQRO, MEXU). Previous records of this species for the peninsula area are here referred to Aechmea tillandsioides (Mart. & Schult, f.) Baker var kienastii (E. Morr.) L. B. S.

(Ramírez, I. M. & Carnevali, G. 1999: 185-194)

Aechmea tillandsioides Baker. Family Bromeliaceae.

Photo by: María Alejandra Gutierrez, FLAAR Mesoamerica, Mar. 10, 2020. Livingston, Izabal. Camera: Canon 1D X Mark II. Lens: Canon EF 300mm IS II USM. Settings: 1/500 sec; f/8; ISO 8,000. 21 FLAAR Mesoamérica

It is interesting to note that the only species known to live at the very top of the canopy in full sunlight is Aechmea tillandsioides (Bromeliaceae), which is a tank epiphyte with a built-in water reservoir, and is usually found growing on ant nests. The nutrients derived from the microfauna in the tank reservoir and from the ant nest apparently compensate for its otherwise nutrient-poor position. (Croat 1978: 23)

Diaspore strategy has diverged in two directions in the family, those with superior ovaries having capsular fruits with plumose, wind-dispersed seeds, and those with inferior ovaries having fleshy fruits dispersed by animals, chiefly birds. Among the latter, Aechmea magdalenae is eaten by coatis (Kaufmann, 1962), and A. tillandsioides, often associated with ant nests, is partly dispersed by ants, I believe. (Croat 1978: 227)

Common in the tops of trees in the full sunlight; less common on exposed branches over the edge of the lake. Inflorescences usually begin to open in the early dry season, but individual plants

Aechmea tillandsioides. Family Bromeliaceae. may flower over a long period, perhaps for “Piñuela” is one of the common names given in the full flowering season. Flowering from some areas of Guatemala. at least January to August. Mature fruits Photo by: Nicholas Hellmuth, FLAAR Mesoamerica, Nov. 25, 2018. Plaza G, Yaxha, Petén, Guatemala. have been seen from May to December.

Camera: Nikon D5. Lens: Nikon AF-S Micro NIKKOR (Croat 1978: 231) 60mm G ED. Settings: 1/60 sec; f/14; ISO 6,400. Río Dulce, Chocón Machacas.

Photo by: Vivian Díaz, FLAAR Mesoamerica, Feb. 14, 2020. Canyon Rio Dulce, a kilometer east of Chocon Machacas, Livingston. Camera: Google Pixel 3 XL. 23 FLAAR Mesoamérica

Aechmea tillandsioides Baker.

Photo by: David Arrivillaga, FLAAR Mesoamerica, Feb. 14, 2020. Biotopo Chocón Machacas, Municipio de Livingston. Camera: Sony Alpha A9 II. Lens: Sony FE 90mm Macro G OSS. Settings: 1/250 sec; f/9; ISO 640. 24 FLAAR Mesoamérica WHERE DID FLAAR MESOAMERICA TEAM FIND AECHMEA TILLANDSIOIDES IN IZABAL?

At Biotope Chocón Machacas I found Aechmea tillandsioides a few meters from the open area after you get up the hill to the level where the original housing area was. A branch had fallen down and the bromeliad was in full bloom on our first visit in February. So, in March I returned to see whether it was still flowering, and yes, it was still flowering!

We also found this species at least once a day, high in trees, almost every area we visited: Lagunita Creek, Canyon Río Dulce, and along the shore of the inlets on both sides of El Golfete. There are so many ecosystems to study in these areas that we have not yet doing fieldwork and photography in the west half of El Golfete nor the western end of the Municipio (west of the highway bridge over Río Dulce).

Landscape of Municipio de Livingston.

Photo by: María Alejandra Gutiérrez, FLAAR Mesoamerica, Feb. 14, 2020. Canyon Rio Dulce, a kilometer east of Chocon Machacas, Livingston, Izabal. Camera: Sony RX10 IV. Lens: Sony FE Macro. Settings: 1/4,000 sec; f/2.8; ISO 500. Aechmea tillandsioides. Family Bromeliaceae. “Piñuela” is one of the common names given in some areas of Guatemala.

Photo by: David Arrivillaga, FLAAR Mesoamerica, Feb. 14, 2020. Biotopo Chocón Machacas, Municipio de Livingston. Camera: Sony Alpha A9 II. Lens: Sony FE 90mm Macro G OSS. Settings: 1/250 sec; f/9; ISO 640. 26 FLAAR Mesoamérica WHERE ELSE HAS AECHMEA TILLANDSIOIDES BEEN FOUND IN IZABAL?

• Missouri Botanical Garden (MO:MO) • Catalog #: 3459083 • Taxon: Aechmea tillandsioides (Mart. ex Schult. & Schult. f.) Baker • Family: Bromeliaceae • Determiner: J. Bitther, • Collector: Rafael Molina 4856 • Date: 1976-10-10 • Locality: Guatemala, Izabal, El Estor, Canaboncito

• Missouri Botanical Garden • Catalog #: 3594981 • Taxon: Aechmea tillandsioides (Mart. ex Schult. & Schult. f.) Baker • Family: Bromeliaceae • Determiner: J. Utley (NOLS), • Collector: W.E. Harmon & J.A. • Fuentes 2117 • Date: 1970-03-14 • Locality: Guatemala, Izabal, 15.8 km S of Modesto Méndez. Collec- tion from along newly constructed road in mixed tropical rainforest. Epiphytes from an oak tree felled by road construction 15.76 -89.1 • Elevation: 70 meters (230ft) Nicholas preparing his photography equipment.

Photo by: Victor Mendoza, FLAAR Mesoamerica, Feb. 14, 2020. Biotopo Chocón Machacas, Livingston, Izabal.

Camera: Google Pixel 3 XL. 27 FLAAR Mesoamérica

WHERE ELSE HAS A. WHERE ELSE HAS A. TILLANDSIOIDES BEEN FOUND IN TILLANDSIOIDES BEEN FOUND IN PETÉN, NOT FAR FROM IZABAL? ALTA VERAPAZ, NEAR IZABAL?

All universities are closed during the So far neither MOBOT herbarium unfortunate Coronavirus pandemic, so we records nor www.naturalista.mx/taxa/283296- have not yet been able to have our team of Aechmea-tillandsioides has any list of botany students visit the helpful herbaria Aechmea tillandsioides for Alta Verapaz. at the universities in Guatemala City.

Aechmea tillandsioides Baker. This is the curious fruit of A. tillandsioides. On this stage, the new fruits have this blue-violet color.

Photo by: David Arrivillaga, FLAAR Mesoamerica, Jul. 9, 2019. Yaxha, Nakum and Naranjo Park, Petén. Camera: Nikon D5. Lens: Nikon AF-S Micro NIKKOR 60mm G ED. Settings: 1/200 sec; f/4.5; ISO 2,000. Aechmea tillandsioides. Family Bromeliaceae. “Piñuela” is one of the common names given in some areas of Guatemala.

Photo by: María Alejandra Gutiérrez, FLAAR Mesoamerica, Mar. 10, 2020. Livingston. Camera: Canon 1D X Mark II. Lens: Canon EF 300mm IS II USM. Settings: 1/250 sec; f/8; ISO 8,000. 29 FLAAR Mesoamérica WHERE ELSE HAS AECHMEA TILLANDSIOIDES BEEN FOUND IN GUATEMALA?

So far we have not yet found the records for collections throughout the rest of Petén. But surely, since this bromeliad is found in Chiapas, Tabasco, southern Campeche, and South- ern Quintana Roo, it will be findable in more areas of Petén besides just Tikal, Yaxha, and Izabal. We need to hike to the multiple herbaria at universities in Guatemala and check where their botanists have collected Aechmea tillandsioides.

Androlepsis skinneri and Aechmea tillandsioides.

Photo by: Senaida Ba, FLAAR Mesoamerica, Nov. 10, 2020. El Golfete, Livingston. Camera: Nikon D5. Lens: Nikon AF-S VR Micro-NIKKOR 105mm IF-ED. Settings: 1/320 sec; f/8; ISO 640. Aechmea tillandsioides Baker. Family Bromeliaceae.

Photo by: Nicholas Hellmuth, FLAAR Mesoamerica, Nov. 8, 2020. Río Lámpara, Livingston. Camera: Nikon D810. Lens: Nikon 200mm AF-D Tele-Macro. Settings: 1/320 sec; f/8; ISO 1,600. 31 FLAAR Mesoamérica AECHMEA TILLANDSIOIDES IN CHIAPAS

Since there are not many local uses of Aechmea tillandsioides, I can understand that Cook does not include Aechmea tillandsioides in her index. However, in , the roots of a mature Aechmea tillandsioides, can be made into a brush to utilize for household chores (Balick and Arvigo 2015: 135). It would help if ethnographers can recheck in their areas of field work.

Aechmea tillandsioides. Aechmea tillandsioides.

Photo by: Nicholas Hellmuth, FLAAR Mesoamerica, Photo by: David Arrivillaga, FLAAR Mesoamerica, Nov. 8, 2020. Río Lámpara, Livingston. Mar, 11. 2020. Livingston.

Camera: Nikon D810. Lens: Nikon Camera: Nikon D810. Lens: Nikon 200mm AF-D Tele-Macro. 200mm AF-D Tele-Macro. Settings: 1/320 sec; f/8; ISO 1,600. Settings: 1/250 sec; f/10; ISO 3,200. 32 FLAAR Mesoamérica

Most plants that are found in Chiapas, Tabasco or Campeche are also in adjacent Petén. Aechmea tillandsioides is present in Chiapas:

www.naturalista.mx/taxa/283296-Aechmea- tillandsioides shows in three places in Chiapas (all three near the border of Guatemala). So, it should be possible to find in northern or northwestern Huehuetenango (which borders on Chiapas) and western Petén (which borders on Chiapas) and north (western) Petén which is just a few miles from where has been found in Campeche.

Aechmea tillandsioides is in for Yaxchilan Natural Monument (Meave et al. 2008: 66). Since Yaxchilan is on one side of the Río Usumacinta, surely Aechmea tillandsioides can be found on the Petén side.

If found at Yaxchilan, no surprise that Aechmea tillandsioides is also documented for Bonampak (SEMARNAT n.d.: 22). And if found in both these areas, nice that it has also been documented for Palenque area (Gomez et al. 2015: 576).

Dr Hellmuth always recommend different equipment to take the best photos on a field trip.

Photo by: Vivian Díaz, FLAAR Mesoamerica, Feb. 14, 2020. Biotopo Chocón Machacas, Livingston, Izabal.

Camera: Google Pixel 3 XL. Aechmea tillandsioides. Family Bromeliaceae. “Piñuela” is one of the common names given in some areas of Guatemala.

Photo by: María Alejandra Gutierrez, FLAAR Mesoamerica, Feb. 13, 2020. Chocón Machacas, Livingston. Camera: Sony RX10 IV. Lens: Sony FE 200-600mm G OSS. Settings: 1/640 sec; f/4; ISO 800. 34 FLAAR Mesoamérica AECHMEA TILLANDSIOIDES IN TABASCO

So far nothing on the map of www.naturalista.mx. But, another web site does list for Tabasco: Campeche, Quintana Roo, Tabasco, Chiapas (so all areas of Mexico that border on Peten, Guatemala).

The same web site also lists Aechmea tillandsioides for Oaxaca and Veracruz.

Platyaechmea tillandsioides is listed for Tabasco (Bueno et al. 2005: 104, without saying where). Platyaechmea tillandsioides is one old name for Aechmea tillandsioides. When you really want to find where Aechmea tillandsioides can be found, it helps to also search for the synonyms.

Aechmea tillandsioides Baker. This is the curious fruit of A. tillandsioides. On this stage, the new fruits have this blue-violet color.

Photo by: David Arrivillaga, FLAAR Mesoamerica, Jul. 9, 2019. Yaxha, Nakum and Naranjo Park, Petén. Camera: Nikon D5. Lens: Nikon AF-S Micro NIKKOR 60mm G ED. Settings: 1/800 sec; f/5.6; ISO 2,000. 35 FLAAR Mesoamérica AECHMEA TILLANDSIOIDES IN CAMPECHE

Mentioned for Campeche in several publications including by Ramírez (2010).

One location in Campeche on the map of www.naturalista.mx.

Aechmea tillandsioides. Family Bromeliaceae. This is the curious fruit of A. tillandsioides. On this stage, the new fruits have this blue-violet color. Photo by: David Arrivillaga, FLAAR Mesoamerica, Jul. 9, 2019. Yaxha, Nakum and Naranjo Park, Petén. Camera: Nikon D810. Lens: Nikon AF-S Micro NIKKOR 60mm G ED. Settings: 1/20 sec; f/11; ISO 2,000. 36 FLAAR Mesoamérica AECHMEA TILLANDSIOIDES IN QUINTANA ROO

One location on the map of www.naturalista.mx. Also:

Nombre científico Colector (es) Aechmea tillandsioides var. tillandsioides E. Cabrera y R. Durán Número de colecta Determinador (es) 4668 Ivón Ramírez Fecha de colecta País 1983-05-07 México (MX) Forma de vida o estadio Estado Epífita Quintana Roo Dependencia Municipio Departamento de Botánica, Felipe Carrillo Puerto Instituto de Biología (IBUNAM)

Localidad Institución A 1 km al este de Chanca Veracruz Universidad Nacional Autónoma de México (UNAM)

Aechmea bracteata Griseb.

Photo by: Erick Flores, FLAAR Mesoamerica, Sep. 5, 2018. Yaxha, Petén. Camera: Canon 1D X Mark II. Lens: Canon EF 300mm IS II USM. Settings: 1/1,100 sec; f/3.2; ISO 1,000. 37 FLAAR Mesoamérica AECHMEA TILLANDSIOIDES IN BELIZE

Three locations for Belize are on the map of www.naturalista.mx.

Plus mention by: (Balick, Nee and Atha 2000: 174). (Balick and Arvigo 2015: 135).

Surely more locations can be found for Belize. Any energetic student could find mention of Aechmea tillandsioides in REA assessments of nature preserves throughout Belize. For example, REA on Flora of the Sarstoon Temash National Park and Surroundings lists Aechmea tillandsioides (Meerman, Herrera, and Howe 2003: Sheet 3 and Sheet 4).

Aechmea tillandsioides.

Photo by: David Arrivillaga, FLAAR Mesoamerica, Feb. 14, 2020. Biotopo Chocón Machacas, Municipio de Livingston. Camera: Sony Alpha A9 II. Lens: Sony FE 90mm Macro G OSS. Settings: 1/250 sec; f/9; ISO 640. Aechmea tillandsioides. Family Bromeliaceae. “Piñuela” is one of the common names given in some areas of Guatemala.

Photo by: David Arrivillaga, FLAAR Mesoamerica, Feb. 14, 2020. Biotopo Chocón Machacas, Municipio de Livingston. Camera: Sony Alpha A9 II. Lens: Sony FE 90mm Macro G OSS. Settings: 1/250 sec; f/10; ISO 1,250. 39 FLAAR Mesoamérica

IN WHAT ECOSYSTEMS CAN YOU FIND NATIVE AECHMEA TILLANDSIOIDES?

We found Aechmea tillandsioides on trees branches in very humid areas (Biotopo Chocón Machacas). Also, in dry- seasonally moist areas (Parque Nacional Yaxhá, Nakum and Naranjo, Petén).

We are curious why no examples are yet documented for adjacent Alta Verapaz, though surely when we ask our plant scouts there and we are allowed to check herbaria lists in Guatemala universities, we will find Aechmea tillandsioides in Alta Verapaz.

DOES AECHMEA TILLANDSIOIDES ALSO GROW IN HOME GARDENS?

This is a popular garden plant sold around the world. So it is found in gardens. But so far we have not noticed this bromeliad in Mayan home gardens.

Nicholas taking photos at Biotopo Chocón Machacas.

Photo by: Juan Pablo Fumagalli, FLAAR Mesoamerica, Mar. 11, 2020. Biotopo Chocón Machacas, Livingston, Izabal.

Camera: Google Pixel 3 XL. Aechmea tillandsioides Baker.

Photo by: Nicholas Hellmuth, FLAAR Mesoamerica, Nov. 8, 2020. Río Lámpara, Livingston. Camera: Nikon D810. Lens: Nikon 200mm AF-D Tele-Macro. Settings: 1/320 sec; f/8; ISO 1,600. 41 FLAAR Mesoamérica USES OF AECHMEA TILLANDSIOIDES

The roots of a mature Aechmea tillandsioides can be made into a brush to utilize for house- hold chores (Balick and Arvigo 2015: 135). This is helpful information especially to avoid using plastic. But a brush is best made from leaves of palm trees because if you tear up the roots then you exterminate this plant.

POTENTIAL MEDICINAL USAGE BY LOCAL PEOPLE

We have not yet found peer-reviewed journal articles on medicinal usage of Aechmea tillandsioides by local people.

Aechmea tillandsioides. Family Bromeliaceae. “Piñuela” is one of the common names given in some areas of Guatemala.

Photo by: Victor Castillo, FLAAR Mesoamerica, Nov. 7, 2020. Río Chocón Machacas, Livingston, Izabal. Camera: Sony Alpha A9 II. Lens: Sony FE 200-600mm G OSS. Settings: 1/640 sec; f/6.3; ISO 2,000. 42 FLAAR Mesoamérica

Aechmea tillandsioides Baker. Family Bromeliaceae.

Photo by: Nicholas Hellmuth, FLAAR Mesoamerica, Nov. 25, 2018. Plaza G, Yaxha, Petén. Camera: Nikon D5. Lens: Nikon AF-S Micro NIKKOR 60mm G ED. Settings: 1/60 sec; f/14; ISO 2,560. 43 FLAAR Mesoamérica

WHEN DOES AECHMEA TILLANDSIOIDES FLOWER?

In México is listed as flowering in April and May.

In Izabal it was flowering in February and March (and surely the flowering will continue into April). I bet it began to flower in January.

Aechmea tillandsioides

Photo by: Nicholas Hellmuth, FLAAR Mesoamérica, Jul. 9, 2019. Naranjo, Yaxha, Nakúm y Naranjo. Camera: iPhone XS. 44 FLAAR Mesoamérica

WHAT ARE THE PRIMARY POLLINATORS OF AECHMEA TILLANDSIOIDES FLOWERS?

Bimodal pollinator by birds and insects, commonly bees.

(Aguilar-Rodríguez, Kromer, Tschapka, García-Franco, Escobedo-Sarti, MacSwiney G 2019: 4)

“Standley considered Codonanthe calcarata Hanst. a synonym of this species, but the name was misapplied according to Leeuwenberg (1958). Like most other members of the genus, C. crassifolia is usually associated with ant nests, though many seedlings are found in cracks in tree bark and are not attended by ants. The ant nests frequently also contain Aechmea tillandsioides (22. Bromeliaceae).”

(Croat 1978: 641)

Fruits are no doubt chiefly dispersed by birds. The seeds ooze from the base when the fruit is pressed in the midsection, in the manner of Anthurium (21. Araceae). Part of the seeds may stick to the bird’s beak, owing to the slender mucilaginous funicles.

(Croat 1978: 232)

Aechmea tillandsioides

Photo by: David Arrivillaga, FLAAR Mesoamerica, Mar, 11. 2020. Livingston, Izabal.

Camera: Nikon D810. Lens: Nikon 200mm AF-D Tele-Macro. Settings: 1/250 sec; f/14; ISO 3,200. Aechmea tillandsioides Baker. Aechmea tillandsioides. Family Bromeliaceae. “Piñuela” is one of the common names given in some areas of Guatemala Photo by: Nicholas Hellmuth, FLAAR Mesoamerica, Jul. 9, 2019. Naranjo, Yaxha, Petén. Photo by: Nicholas Hellmuth, FLAAR Mesoamerica, Nov. 8, Camera: iPhone XS. 2020. Río Lámpara, Livingston. Camera: Nikon D810. Lens: Nikon 200mm AF-D Tele-Macro. Settings: 1/32o sec; f/8; ISO 1,600. 46 FLAAR Mesoamérica CONCLUDING DISCUSSION AND SUMMARY ON AECHMEA TILLANDSIOIDES

I did not learn about the relationship of ants with the roots of until I was back in my office, 300 kilometers away from these bromeliads in Izabal areas. So definitely will inspect the roots the next time we are hiking through the nature reserves of Municipio de Livingston and boating through the Río Dulce Canyon.

It will also help on a future field trip to see whether Aechmea tillandsioides bromeliads are still flowering in months other than February and March.

Our long-range goal is to make a map of Municipio de Livingston, and have one map for orchids, one map for bromeliads, etc. to show where you can see and photograph these plants when you arrive here yourself.

Aechmea tillandsioides Baker. Family Bromeliaceae.

Photo by: Senaida Ba, FLAAR Mesoamerica, Nov. 5, 2020. Mirador del Cañón, Río Dulce, Livingston. Camera: Nikon D5. Lens: Nikon AF-S VR Micro-NIKKOR 105mm IF-ED. Settings: 1/125 sec; f/6.3; ISO 2,560. Aechmea tillandsioides.

Photo by: Nicholas Hellmuth, FLAAR Mesoamerica, Nov. 10, 2020. El Golfete, Río Negro, Livingston. Camera: Nikon D810. Lens: Nikon 200mm AF-D Tele-Macro. Settings: 1/250 sec; f/9; ISO 4,000. 48 FLAAR Mesoamérica

BIBLIOGRAPHY, REFERENCES CITED AND SUGGESTED READING ON AECHMEA TILLANDSIOIDES

Most helpful monographs on this plant

Croat of MOBOT has published helpful information (1978). It is especially helpful that this report is available on-line as an easy download.

SMITH, Lyman B. and Robert Jack DOWNS 1974 Pitcairnioideae (Bromeliaceae). Flora Neotropica. Vol. 14, No. 1, (Oct. 14, 1974), pp. 1-609+611-641+643-658. New York Botanical Garden Press on behalf of Organization for Flora Neotropica

If this is available as a digital edition, open (so you can search and so you can copy-and- paste to cite) we will use it. Otherwise Standley and Steyermark and Standley and Williams are the best sources, especially since Standley and also Williams were interested in ethno- botanical information; after they passed away the subsequent botanical monographs did not include information on how local Maya and other local people used the plants.

Most helpful articles on this plant

Need to keep looking to find an article on Aechmea tillandsioides of Mesoamérica.

Most helpful web sites on this plant

Need to accomplish lots more research to find a website with helpful info on a single species. Most websites on bromeliads are for garden varieties; we are studying wild native bromeliads. 49 FLAAR Mesoamérica

AGUILAR-Rodríguez, Pedro, KRÖMER, Thorsten, TSCHAPKA, Marco, GARCÍA-Franco, José, ESCOBEDO-Sarti, Jeanett and McSWINEY G., Cristina 2019 Bat pollination in Bromeliaceae. Plant Ecology & Diversity Journal. Taylor & Francis Group. Volume 12. Pages 1-19

ATRAN, Scott, LOIS, Mimena and Edilberto UCAN Ek’ 2004 Plants of the Petén Itza’ Maya. Museum of Anthropology, Memoirs, Number 38, University of Michigan. 248 pages. Very helpful and nice collaboration with local Itza’ Maya people. But would help in the future to have a single index that has all Latin, Spanish, and English plant names so that you can find plants more easily.

Not available as a download.

BALICK, Michael J., NEE, Michael H. and Daniel E. ATHA 2000 Checklist of the Vascular Plants of Belize: With Common Names and Uses. Memoirs of the New York Botanical Garden Vol. 85. 246 pages.

BALICK, Michael J. and Rosita ARVIGO 2015 Messages from the Gods: A Guide to the Useful Plants of Belize. The New York Botanical Garden, Oxford University Press.

BUENO, Joaquín, ALVAREZ, Fernando and Silvia SANTIAGO (Editors) 2005 Biodiversidad del Estado de Tabasco. CONABIO, Instituto de Biologia, UNAM, Mexico. 370 pages

COOK, Suzanne 2016 The forest of the Lacandon Maya: an ethnobotanical guide. Springer. 334 pages.

One of the most helpful and informative monographs on ethnobotanical aspects of plants of this part of the Lacandon area of Chiapas.

Probably since Aechmea tillandsioides is nowadays not often used by local people, this colorful bromeliad is not in her botanical index. However it would be worth checking all Lacandon and Q’eqchi’ Mayan areas of Chiapas and Guatemala to see if the roots are used, since for Belize this is a use listed by Balick and Arvigo.

Sold online: www.springer.com/la/book/9781461491101 50 FLAAR Mesoamérica

CROAT, Thomas B. 1978 Flora of Barro Colorado Island. Stanford University Press. 943 pages.

https://books.google.com.gt/books?id=Whc_ahfhFFoC&pg=PA231&lpg= PA231&dq=thomas+b.+croat+aechmea+tillandsioides&source=bl&ots=VqnS B1TtWt&sig=ACfU3U13QTQZnzzBqTI1e4SgbYytSEj_qw&hl=en&sa =X&ved=2ahUKEwjOv4ikrffsAhUDqlkKHQPGD2MQ6AEwDnoECAgQAg#v =onepage&q=thomas%20b.%20croat%20aechmea%20tillandsioides&f=false

ESPEJO-Serna, Adolfo and Ana Rosa LÓPEZ-Ferrari 2010 La familia Bromeliaceae en México. Botanical Sciences 96 (3): 533-554, 2018.

Downloadable: www.scielo.org.mx/pdf/bs/v96n3/2007-4476-bs-96-03-533.pdf

GÓMEZ-DOMÍNGUEZ, HÉCTOR, PÉREZ FARRERA, MIGUEL ÁNGEL ESPINOZA JIMÉNEZ Josefa ANAHÍ and Mirna Ivett MARQUEZ Reynoso 2015 LISTADO FLORÍSTICO DEL PARQUE NACIONAL PALENQUE, CHIAPAS, MÉXICO. Botanical Sciences 93 (3): 559-578, 2015.

LAUBE, Stefan and Ferhard ZOTZ 2006 Neither Host-specific nor Random: Vascular Epiphytes on Three Tree Species in a Panamanian Lowland Forest. Annals of Botany 97: 1103–1114, 2006.

Downloadable: www.ncbi.nlm.nih.gov/pmc/articles/PMC2803392/pdf/mcl067.pdf

LUNDELL, Cyrus L. 1937 The Vegetation of Peten. Carnegie Institution of Washington, Publ. 478. Washington. 244 pages.

LUNDELL, Cyrus L. 1938 Plants Probably Utilized by the Old Empire Maya of Peten and Adjacent Lowlands. Papers of the Michigan Academy of Sciences, Arts and Letters 24, Part I:37-59.

MARTÍNEZ, J. N 2011 Estado actual y valor de uso etnobotánico de las especies vegetales utilizadas en la industria artesanal alfarera del municipio de Guatajiagua, Morazán El Salvador. Universidad de El Salvador. 54 pages. 51 FLAAR Mesoamérica

MEAVE, Jorge A., ROMERO-Romero, Marco A.,VALLE-DOMÉNECH, ANDRÉS;RINCÓN-GUTIÉRREZ, ARMANDO;MARTÍNEZ, ESTEBAN and Clara H. RAMOS 2008 Plant diversity assessment in the Yaxchilán Natural Monument, Chiapas, México Boletín de la Sociedad Botánica de México, Núm. 83, sin mes, 2008, pp. 53-76. Sociedad Botánica de México, A.C. México.

Downloadable from Redalyc.

MEERMAN, J. C., HERRERA, P. and A. HOWE 2003 Rapid Ecological Assessment Sarstoon Temash National Park, Toledo District, Belize Volume II: Appendices (Species lists and raw data). Sarstoon Temash Institute for Indigenous Management (SATIIM).

Downloadable on the Internet.

MONDRAGÓN-Chaparro, Demetria Martha, RAMÍREZ-Morillo, Ivón Mercedes, FLORES Cruz, María and José Guadalupe GARCÍA-Franco 2011 La familia Bromeliaceae en México. Universidad Autónoma Chapingo. Grupo Publicitario Imagen Digital, Texcoco, Edo. de México

MORALES-Linares, Jonas Spatial structure of ant-gardens: Vertical distribution on host trees and succession/ segregation of associated vascular epiphytes.

www.researchgate.net/publication/318201906_Spatial_structure_of_ant-gardens_ Vertical_distribution_on_host_trees_and_successionsegregation_of_associated_ vascular_epiphytes/link/59b06c42458515a5b484e061/download

RAMÍREZ Morillo, Ivón M. and F. C. GERMÁN Carnevali 1999 A new species of Tillandsia, some new records and a checklist of the Bromelliaceae from the Yucatán Peninsula. Harvard Papers in Botany Vol. 4, No. 1 (September 1999), pp. 185-194.

RAMÍREZ Morillo, Ivón M. 2010 Bromelias. pp 228-233 In; La Biodiversidad en Campeche: Estudio de Estado. Comisión Nacional para el Conocimiento y Uso de la Biodiversidad (conabio), Gobierno del Estado de Campeche, Universidad Autónoma de Campeche, El Colegio de la Frontera Sur. México. 730 pages. Volume coordinators: Villalobos- Zapata, G. J., and J. Mendoza Vega (Coord.)

Download: http://bioteca.biodiversidad.gob.mx/janium/Documentos/7371.pdf 52 FLAAR Mesoamérica

SELVEN Pérez, Edgar and Miriam Lorena CASTILLO Villeda 2000 A rapid assessment of avifaunal diversity in aquatic habitats of Laguna del Tigre National Park, Petén, Guatemala. Pages 56-60 in: A Biological Assessment of Laguna del Tigre National Park, Petén, Guatemala, Bestelmeyer, B.T. and Alonso, L.E. (editors). Conservation International.

SEMARNAT n.d. Programa de Conservación y Manejo Monumento Natural Bonampak. SEMARNAT, México.

SMITH, Lyman B. and Robert Jack DOWNS 1974 Pitcairnioideae (Bromeliaceae). Flora Neotropica. Vol. 14, No. 1, (Oct. 14, 1974), pp. 1-609+611-641+643-658. New York Botanical Garden Press on behalf of Organization for Flora Neotropica

STANDLEY, Paul C. and Samuel J. RECORD 1936 The Forests and Flora of British . Field Museum of Natural History. Publication 350, Botanical Series Volume XII. 432 pages plus photographs.

STANDLEY, Paul C. and Julian A. STEYERMARK 1958 Flora of Guatemala. Fieldiana: Botany, Vol. 24, Part I. Chicago Natural History Museum.

Downloadable from several web sites. Here is one:

https://libsysdigi.library.illinois.edu/oca/Books2009-04/floraofguatemala1fistan/ floraofguatemala1fistan.pdf

VÉLIZ Mario, MIJANGOS, Ismael and Vilma FIALKO 2010 Guía florística del sitio arqueológico Naranjo-Saal, Petén, Guatemala. IDAEH-WMF- PIAREN-FCG, 31 pages.

Nicely designed, helpful, list of orchids and bromeliads of the Naranjo site area of Parque Nacional Yaxha Nakum Naranjo. Archaeologist Vilma Fialko also saves all orchids and bromeliads which are found blown over in heavy rain storms.

VOGL, C. R., VOGL-Lukasser, B. and J. CABALLERO 2002 Homegardens of Maya Migrants in the District of Palenque (Chiapas/México): Implications for Sustainable Rural Development. In: Stepp, J.R., Wyndham, F.S., and R.K. Zarger (eds.). Ethnobiology and Biocultural Diversity. Pp: 631 – 647. University of Georgia Press; Athens, Georgia 53 FLAAR Mesoamérica

LEE, Jones 1972 A Highland Maya People and their Habitat: The Natural History, Demography and Economy K´ekchi´ PhD dissertation. 475 pages.

His field work was near San Pedro Carcha, which is now a suburb of Cobán, Alta Verapaz. The climate is moist due to moist clouds during many times of the year.

Downloadable on the Internet.

WOLF, J. H. D. and A. FLAENCO S. 2003 Patterns in species richness and distribution of vascular epiphytes in Chiapas, México. J. Biogeogr. 30(11): 1689-1707. 54 FLAAR Mesoamérica

HELPFUL WEB SITES FOR ANY AND ALL PLANTS

There are several web sites that are helpful even though not of a university or botanical garden or government institute.

However most popular web sites are copy-and-paste (a polite way of saying that their authors do not work out in the field, or even in a botanical garden). Many of these web sites are click bait (they make money when you buy stuff in the advertisements that are all along the sides and in wide banners also. We prefer to focus on web sites that have reliable information.

https://serv.biokic.asu.edu/neotrop/plantae/ Neotropical Flora data base. To start your search click on this page: https://serv.biokic.asu.edu/neotrop/plantae/collections/harvestparams.php

http://enciclovida.mx CONABIO. The video they show on their home page shows a wide range of flowers pollinators, a snake and animals. The videos of the insects are great.

www.kew.org/science/tropamerica/imagedatabase/index.html Kew gardens in the UK is one of several botanical gardens that I have visited (also New York Botanical Gardens and Missouri Botanical Gardens (MOBOT), in St Louis. Also the botanical garden in Singapore and El Jardín Botánico, the open forest botanical garden in Guatemala City).

www.ThePlantList.org This is the most reliable botanical web site to find synonyms. In the recent year, only one plant had more synonyms on another botanical web site. 55 FLAAR Mesoamérica

WEB SITES SPECIFICALLY ON AECHMEA GENUS OR AECHMEA TILLANDSIOIDES SPECIES

www.cicy.mx/sitios/flora%20digital/ficha_virtual.php?especie=173 Often a helpful web site but very little information on Aechmea tillandsioides. But does at least mention that it is found in Tabasco, Campeche and Quintana Roo. However it is only rarely mentioned for Tabasco, but considered common in habitats that are appropriate (southern Campeche and southern Quintana Roo).

http://www.fcbs.org/pictures/Aechmea.htm Lists all species of Aechmea (but does not indicate where they are found).

www.maya-ethnobotany.org This web site of FLAAR (USA) and FLAAR Mesoamerica (Guatemala) has two pages on this plant: one where found in Parque Nacional Yaxha, Nakum and Naranjo by our team last year. And one page where found in Biotopo Chocon Machacas, Municipio de Livingston, Izabal, Guatemala. We estimate this species is very common on trees throughout the Municipio de Livingston.

www.naturalista.mx/taxa/283296-Aechmea-tillandsioides Gorgeous photos (though the color balance may be a tad off; the red is a bit too red). 56 FLAAR Mesoamérica APPENDIX A

Where has Aechmea tillandsioides been stored in Botanical Herbaria that are listed on the Neotropical Flora data base

• Missouri Botanical Garden • Catalog #: 3459083 • Taxon: Aechmea tillandsioides (Mart. ex Schult. & Schult. f.) Baker • Family: Bromeliaceae • Determiner: J. Bitther, • Collector: Rafael Molina 4856 • Date: 1976-10-10 • Locality: Guatemala, Izabal, El Estor, Canaboncito

• Missouri Botanical Garden • Catalog #: 3594981 • Taxon: Aechmea tillandsioides (Mart. ex Schult. & Schult. f.) Baker • Family: Bromeliaceae • Determiner: J. Utley (NOLS), • Collector: W.E. Harmon & J.A. Fuentes 2117 • Date: 1970-03-14 • Locality: Guatemala, Izabal, 15.8 km S of Modesto Méndez. Collection from along newly constructed road in mixed tropical rainforest. Epiphytes from an oak tree feld by road construction 15.76 -89.1 • Elevation: 70 meters (230ft) 57 FLAAR Mesoamérica

• Missouri Botanical Garden • Catalog #: 1976048 • Taxon: Aechmea tillandsioides (Mart. ex Schult. & Schult. f.) Baker • Family: Bromeliaceae • Determiner: J. Utley • Collector: Thomas B. Croat 24678 • Date: 1973-06-18 • Locality: Guatemala, Petén, Vicinity of Archeological camp on north shore of Lake Yaxha. Disturbed forest 17.06 -89.39 • Elevation: 152 meters (499ft)

• Missouri Botanical Garden • Catalog #: 100276492 • Taxon: Aechmea tillandsioides (Mart. ex Schult. & Schult. f.) Baker • Family: Bromeliaceae • Determiner: M. Véliz • Collector: 1265 • Date: 1989-03-10 • Locality: Guatemala, Petén, Yaxhaw, El Petén.

• Missouri Botanical Garden • Catalog #: 1793184 • Taxon: Aechmea tillandsioides var. kienastii (E. Morren ex Mez) L.B. Sm. • Family: Bromeliaceae • Determiner: L.B. Smith, • Collector: C.L. Lundell 15498 • Date: 1959-01-29 • Locality: Guatemala, Petén, Tikal National Park, Tikal. In ramonal covering the ruins 17.23 -89.61 • 58 FLAAR Mesoamérica APPENDIX B

Where has Aechmea tillandsioides been located in Guatemala Based on herbaria of Guatemala (which are not yet digitized)

We will include this information in a second edition (when the herbaria are open).

APPENDIX C

Ant Relationships with Aechmea tillandsioides

While doing research on Aechmea tillandsioides I found several articles and reports which mentioned a relationship of ants and Aechmea tillandsioides. Since these ants are evidently focused on the roots, I have not yet noticed them because the flowers are so photogenic, I must admit I focused totally on the flowers. We do not yet have our own documentation nor photos of ants associated with the roots of Aechmea tillandsioides in Municipio Livingston. Nonetheless, we show here an introductory bibliography on this ant relationship.

We are long ago familiar with the well-known ant symbiosis with bull-horn acacia (“subin” is the local name for these kinds of plants). And of course we are equally familiar with the ant symbiosis of the hollow trunks of Cecropia trees (“guarumo” is the local name for these trees).

Plus about two years ago, while hiking along trails looking for wild vanilla orchid vines in really remote areas south of Laguna Lachua (Alta Verapaz), we noticed that most of the Costus flowers had ants on them. Same in the Biotopo Chocón Machacas: at least 40% of theCostus flowers had ants on them. In other words, we enjoy studying, learning about, photographing, and then publishing about ant relationships with plants in Guatemala.

MORALES-Linares, Jonas Spatial structure of ant-gardens: Vertical distribution on host trees and succession/ segregation of associated vascular epiphytes.

www.researchgate.net/publication/318201906_Spatial_structure_of_ant-gardens_Vertical_distribution_on_host_trees_ andsuccessionsegregation_of_associated_vascular_epiphytes/link/59b06c42458515a5b484e061/download 620000 650000 680000 710000 740000

BELICE Livingston, 1780000 Mar Caribe Izabal N

NE NW

Petén E

W

3. SE

Sarstoon River SW S

Modesto Río Sarstún Bahía de 1760000 Mendez CA13 Amatique

Municipio de Livingston Livingston Municipio de CA13 2. Livingston CA13 Municipio de El Golfete

1740000 Puerto Barrios Puente de Puerto CA13 Río Dulce Municipio de Barrios KM 274 Livingston CA13 Municipio de CA9 El Estor 1720000 Lago Izabal La Ruidosa Áreas naturales CA9 protegidas de Izabal 1. Municipio de Morales 1. Bocas del Polochic (El Estor) 1700000 CA9 2. Punta de Manabique (Puerto Barrios)

3. Sarstoon Temash Municipio de National Park Los Amates (Belice) HONDURAS

1680000 Áreas naturales Zacapa Hacia Ciudad de protegidas de Livingston Guatemala

Petén BELICE Bahía de 4. Amatique 1760000 4.

6. Municipio de Livingston El Golfete 5. 2. 3.

7. 1740000

2.

Municipio de Municipio de Puerto Barrios El Estor 1. Municipio de 1.

Lago Izabal Morales 1720000

620000 640000 660000 680000 700000

Izabal 1. Área sin protección 4. Área de 6. Biotopo Protegido Usos Múltiples Chocón Machacas 2. Parque Nacional Río Sarstún Río Dulce 5. Sierra de 7. Reserva Protectora 3. El Higuerito Santa Cruz de Manantiales Cerro San Gil Elaborado por: Andrea de la Paz; Amanda Estrada Rodas. FLAAR Mesoamerica 2020 60

ACKNOWLEDGEMENTSACKNOWLEDGEMENTS TO FLAAR MESOAMÉRICA TO FLAAR MESOAMÉRICA

Flor de Maria Setina is the office manager, overseeing all Juan Carlos Hernandez takes the material that we write the diverse projectsThe reportsaround theare worlda joint (including production FLAAR- between theand field places trip it intoteam the and pertinent the in-house modern Internet software to REPORTS research on advanced officewide-format team. digital So here inkjet we wish toproduce cite the our full web team: pages (total network is read by over half a printers, a worldwide project for over 20 years). We also million people around the world). utilize the inkjet prints to produce educational banners to Flor de María Setina is the office manager, overseeing all David Arrivillaga is an experienced photographer and donate to schools. Paulo Nuñez is a webmaster, overlooking the multitude the diverse projects around the world (including FLAAR- is able to handle both Nikon and the newest Sony digital of web sites. Internet SEO changes every year, so we work REPORTS research on advanced wide-format digital inkjet cameras. Work during and after a field trip also includes Vivian Díaz environmental engineer, is project manager together to evolve the format of our web sites. printers, a worldwide project for over 20 years). We also sorting, naming, and processing. And then preparing reports for flora, fauna projects (field work and resulting reports utilize the inkjet prints to produce educational banners to in PDF format. at a level helpful for botanists, zoologists and ecologists, Valeria Aviles is an illustrator for MayanToons, the division donate to schools. and for university students). Also coordinates activities at in charge of educational materials for schools, especially the Juan Carlos Hernández takes the material that we write MayanToons, division where educational material for kids is Q’eqchi’ Mayan schools in Alta Verapaz, Q’eqchi’ and Petén Vivian Díaz environmental engineer, is project manager and places it into the pertinent modern Internet software to prepared. Itzá Maya in Petén, and the Q’eqchi’ Mayan and Garifuna for flora, fauna projects (field work and resulting reports produce our web pages (total network is read by over half a schools in the municipality of Livingston, Izabal. at a level helpful for botanists, zoologists and ecologists, million people around the world). Victor Mendoza identifies plants, mushrooms, lichen, and for university students). Also coordinates activities at insects, and arachnids. When his university schedule allows, Josefina Sequen is illustrator for MayanToons and also helps MayanToons, division where educational material for kids is Paulo Núñez is a webmaster, overlooking the multitude he also likes to participate in field trips on flora and fauna prepare illustrations for Social Media posts and for animated prepared. of web sites. Internet SEO changes every year, so we work research. videos. together to evolve the format of our web sites. Victor Mendoza identifies plants, mushrooms, lichen, Vivian Hurtado prepares the bibliography for each subject Rosa Sequen is also an illustrator for MayanToons and also insects, and arachnids. When his university schedule allows, Valeria Avilés is an illustrator for MayanToons, the division and downloads pertinent research material for our e-library helps prepare illustrations for Social Media posts and for he also likes to participate in field trips on flora and fauna in charge of educational materials for schools, especially the on flora and fauna. All of us use both these downloads plus animated videos. research. Q’eqchi’ Mayan schools in Alta Verapaz, Q’eqchi’ and Petén our in-house library on flora and fauna of Mesoamerica Itzá Maya in Petén, and the Q’eqchi’ Mayan and Garifuna (Mexico through Guatemala into Costa Rica). Laura Morales is preparing animated videos in MayanToons Vivian Hurtado prepares the bibliography for each subject schools in the municipality of Livingston, Izabal. style since animated videos are the best way to help and downloads pertinent research material for our e-library Andrea de la Paz is a designer who helps prepare the master- school children how to protect the fragile ecosystems and on flora and fauna. All of us use both these downloads plus Josefina Sequen is illustrator for MayanToons and also helps plan for aspects of our publications. She is our editorial art endangered species our in-house library on flora and fauna of Mesoamerica prepare illustrations for Social Media posts and for animated director (Mexico through Guatemala into Costa Rica). videos. Heidy Alejandra Galindo Setina joined our design team in Senaida Ba is photography assistant for many years. She August 2020. She likes photography, drawing, painting, and design. Andrea de la Paz is a designer who helps prepare the master- Rosa Sequen is also an illustrator for MayanToons and also knows the Canon, Nikon and is learning the two new Sony plan for aspects of our publications. She is our editorial art helps prepare illustrations for Social Media posts and for mirrorless cameras. She prepares, packs, sets-up, and Maria José Rabanales she is part of the team for editing director animated videos. helps the photographers before, during, and after each photographic reports and educational material of Flora day’s field trip. and Fauna since September 2020. She works together with Senaida Ba is photography assistant for many years. She Laura Morales is preparing animated videos in MayanToons others of the team to prepare the finished pdf editions of the knows the Canon, Nikon and is learning the two new Sony style since animated videos are the best way to help Jaqueline Gonzalez is a designer who puts together the material of the Yaxha, Nakum and Naranjo Project. mirrorless cameras. She prepares, packs, sets-up, and school children how to protect the fragile ecosystems and text and photographs to create the actual report (we have helps the photographers before, during, and after each endangered species several designers at work since we have multiple reports Alejandra Valenzuela, biology student is now part of Flora day’s field trip. to produce). y Fauna’s photographic report and educational material Heidy Alejandra Galindo Setina joined our design team in editing team since September 2020. Jaqueline González is a designer who puts together the August 2020. She likes photography, drawing, painting, and design. Roxana Leal is Social Media Manager for flora and fauna text and photographs to create the actual report (we have research and publications, and MayanToons educational Cristina Ríos designer student who join the editorial team several designers at work since we have multiple reports Maria José Rabanales sheis part of the team for editing book projects on December 2020. She will combine the text, pictures and to produce). photographic reports and educational material of Flora maps into the FLAAR Mesoamerica editorial criteria. and Fauna since September 2020. She works together with Maria Alejandra Gutierrez is an experienced photographer, Roxana Leal is Social Media Manager for flora and fauna others of the team to prepare the finished pdf editions of the especially with the Canon EOS 1D X Mark II camera and Alexander Gudiel: designer who join the editorial design research and publications, and MayanToons educational material of the Yaxha, Nakum and Naranjo Project. 5x macro lens for photographing tiny insects, tiny flowers, team on December 2020. He will combine the text, pictures book projects and tiny mushrooms. Work during and after a field trip and maps into the FLAAR Mesoamerica editorial criteria. Alejandra Valenzuela, biology student is now part of Flora also includes sorting, naming, and processing. And then María Alejandra Gutiérrez is an experienced photographer, y Fauna’s photographic report and educational material preparing reports in PDF format. Carlos Marroquín is a USAC graphic design student who especially with the Canon EOS 1D X Mark II camera and editing team since September 2020. volunteered to do his professional practice with the Editorial 5x macro lens for photographing tiny insects, tiny flowers, David Arrivillaga is an experienced photographer and Design team. We are very grateful to people like him who and tiny mushrooms. Work during and after a field trip Alexander Gudiel: designer who joined the editorial design is able to handle both Nikon and the newest Sony digital join our team and bring his knowledge and work. also includes sorting, naming, and processing. And then team on December 2020. He will combine the text, pictures cameras. Work during and after a field trip also includes preparing reports in PDF format. and maps into the FLAAR Mesoamerica editorial criteria. sorting, naming, and processing. And then preparing reports in PDF format. The current Alcalde of Livingston, Mr. Daniel Pinto, together with his team of International Cooperation division, Mr. Edwin Mármol, have set the goal of achieving the municipality development in the years 2020-2024 based on the goals and indicators proposed by the 2030 Agenda for Sustainable Development. From this agenda, FLAAR Mesoamerica will collaborate to achieve Sustainable Development Goal (SDG) number 15 “Life on Land”.

Throughout this cooperation project, different materials have been prepared, like this Photo Essay, that helps to collect information on species, different ecosystems: terrestrial, wetlands and fresh water biodiversity. This information would also be useful as part of a strategy to protect threatened species and prevent their extinction. The municipality’s goals include to promote the sustainable use, conservation and research of the species of flora and fauna of the terrestrial, wetlands and aquatic shore and coastal ecosystems of the Guatemalan Caribbean. Learn more about this project and the SDG indicators at: https://flaar-mesoamerica.org/rain-forests-rivers-lakes-bays-ocean-caves-canyons-livingston-the- caribbean-biodiversity-wonderland-of-guatemala/

SERIES OF MUNICIPIO OF LIVINGSTON

FLAAR Mesoamerica is the creator of the design and authorship of the document. When sharing information or designs on social networks, you must tag the page of FLAAR Mesoamérica, its authors and photographers. In the case of written documents, use the corresponding quote.

Any school, college, university, botanical garden, zoological FLAAR (in USA) and FLAAR Mesoamerica (in Guatemala) garden, botanical or zoological association (or club) may are both non-profit research and educational institutes, so there is no fee. And you do not need to write and post this report on their web sites, (at no cost) as long as ask permission; but we do appreciate when you include they link back to one of our web sites: a link back to one of our sites. Any school, college, university, botanical garden, etc. can post this PDF on www.maya-ethnobotany.org their school or university or institute website for their www.maya-ethnozoology.org students to download at no cost. And you do not need to write and ask permission; but we do appreciate it www.maya-archaeology.org when you include a link back to one of our web sites. www.digital-photography.org www.FLAAR-Mesoamerica.org Any website in or related to the Municipio of Livingston, is also welcome to post this PDF on their web site (no This report may be cited with this information: fee). This permission includes travel agencies, hotels, guide services, etc. And you do not need to write Hellmuth, N. (2020) Bright Red and Yellow, Aechmea and ask permission; but we do appreciate it when you tillandsioides. Biotopo Chocón Machacas Livingston, include a link back to one of our web sites. CECON- Izabal, Guatemala: FLAAR Mesoamerica. USAC, CONAP, FUNDAECO, Plantemos, AIESEC, are welcome to publish our reports, at no cost.

BACK COVER PHOTO All national parks, nature reserves, and comparable are Aechmea tillandsioides. welcome to have and use our reports at no cost. USAC, UVG, URL, Universidad Rural, INTECAP and other Photo by: David Arrivillaga, FLAAR Mesoamerica, Feb. 14, Guatemalan universities, and high schools, and schools, 2020. Biotopo Chocón Machacas, Livingston, Izabal. are welcome to post our reports, at no cost.

Camera: Sony a9 II. Lens: Sony FE 90mm OSS. © Copyright 2020 FLAAR Mesoamerica. Settings: 1/250 sec; f/10; ISO 640. 62 FLAAR Mesoamérica

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